奶山羊IGF-I和IGF-IR基因多态性及其与产奶、体尺性状的相关分析
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摘要
本研究采用PCR-SSCP和DNA测序等技术,检测了关中奶山羊和西农萨能奶山羊2个品种共708个个体的胰岛素样促生长因子-I(IGF-I)基因和其受体基因(IGF-IR)14个基因座的遗传变异,分析了其遗传结构和遗传多样性。同时利用SPSS(16.0)软件,运用最小二乘线性模型等方法对2个奶山羊品种在上述基因座中的多态性与产奶、体尺性状进行相关分析,研究这些基因多态性对奶山羊产奶和生长发育性状的遗传效应,以期发现对我国奶山羊品种重要经济性状具有显著效应的遗传标记,为其高效选育提供遗传学依据。本研究获得了以下结果:
1. 2个奶山羊品种IGF-I基因多态性
本研究共检测了奶山羊IGF-I基因5个基因座(P1～P5),即外显子1W、1、3、4、5以及部分内含子区域。在所研究的5个基因座中,仅P1和P4基因座发现多态性,分别位于外显子1W和第4内含子上。与已公布的山羊IGF-I基因序列(GeneBank登录号:D26119.2)比对发现,该两处多态性是g.1617 G > A和g.5752 G > C点突变导致的,在羊上为首次发现。
卡方检验显示:在P1基因座上,关中奶山羊和西农萨能奶山羊均处于Hardy-Weinberg平衡状态( P>0.05 ) ,其群体遗传多态指数Ne/PIC/He分别为:1.16/0.13/0.14,1.23/0.17/0.19。在P4基因座上,西农萨能奶山羊处于Hardy-Weinberg平衡状态(P>0.05),而关中奶山羊则处于不平衡状态(P<0.01)。关中奶山羊和西农萨能奶山羊群体遗传多态指数Ne/PIC/He分别为:1.43/0.26/0.3,1.72/0.33/0.42。
上述结果表明,P4基因座在这2个奶山羊品种中(关中和西农萨能)多态信息含量中等丰富(0.25 2. IGF-I基因多态性与2个奶山羊品种产奶性状和体尺性状的相关性
在IGF-I基因P1基因座中检测出AG和GG 2种基因型,其中GG基因型在2个品种中都为优势基因型。利用SPSS(16.0)软件,运用最小二乘线性模型等方法分析发现,P1基因座不同基因型与2个奶山羊品种的体尺性状,西农萨能奶山羊第一胎和第二胎产奶量无显著相关(P>0.05)。在分析西农萨能奶山羊奶成分中的脂肪、蛋白质、干物质、非脂固形物、乳糖含量和乳密度6个指标中,AG基因型个体乳脂率显著高于GG型个体(P<0.05),而其乳密度则显著低于GG型个体(P<0.05),不同的基因型对其他4个指标无显著影响。
P4基因座上共检测出CC、CG和GG 3种基因型,其中GG基因型在2个奶山羊品种中是优势基因型。在体尺性状上,关中奶山羊GG基因型仅在胸围上显著大于CG型和CC型(P<0.05),而CG型又显著大于CC型(P<0.05);西农萨能奶山羊群体中,CG基因型个体的体长最大,显著大于CC型和GG型(P<0.05),GG型和CC型差异则不显著(P>0.05)。P4基因座不同基因型对于西农萨能奶山羊产奶性状的影响则表现为:CC基因型个体无论是在第一胎还是第二胎产奶量上都显著高于其他两种基因型(P<0.05),CG基因型个体在第二胎产奶量上又显著高于GG基因型(P<0.05),不同基因型对奶成分无显著影响。
上述结果表明,P1基因座不同基因型与西农萨能奶山羊奶成分中的乳脂率和乳密度存在相关,P4基因座不同基因型与奶山羊体尺性状和产奶量存在相关。
3. 2个奶山羊品种IGF-IR基因多态性
本试验共检测了奶山羊IGF-IR基因9个基因座(R1～R9),包含该基因1～10个外显子及部分内含子(除外显子7)。在这9个基因座上,仅在R8和R9基因座上发现了多态性,2个品种在这2个位点上都表现CC、CT、TT 3种基因型。首次在R8基因座外显子9上发现了C>T的突变(NC_007319:g26688C>T),该突变导致了IGF-IR基因编码的蛋白第608位上的Leu氨基酸发生了同义突变。而R9基因座上的多态性是由于第10内含子上T>C(NC_007319:g28273T>C)突变引起的。
卡方检验显示:在R8和R9基因座中,2个品种均处于Hardy-Weinberg不平衡状态(P<0.01)。在R8中,关中奶山羊和西农萨能奶山羊群体遗传多态指数Ne/PIC/He分别为2.00/0.37/0.50,1.97/0.37/0.50。在R9中则为1.99/0.37/0.50,1.95/0.37/0.50。说明,R8和R9基因座在2个奶山羊品种中多态信息含量为中等丰富(0.25 4. IGF-IR基因多态性与2个奶山羊品种产奶性状和体尺性状的相关
在IGF-IR基因R8基因座中检测出CC、CT、TT 3种基因型,其中CT型在2个品种中都为优势基因型。但是该基因座上不同的基因型对2个奶山羊品种的体尺性状和西农萨能奶山羊的第一胎、第二胎产奶量都不存在着影响(P>0.05)。在所检测的奶成分共6个指标中,CC基因型的西农萨能奶山羊的乳密度显著高于CT基因型(P<0.05),与TT型差异不显著(P>0.05),TT基因型和CT基因型之间差异也不显著(P>0.05)。
在R9基因座中同样检测出CC、CT、TT型3种基因型,其中TT基因型占多数。该基因座不同基因型对西农萨能奶山羊的产奶性状不存在着显著影响。在关中奶山羊中,虽然CC和TT基因型个体的体高分别极显著和显著的高于CT基因型(P<0.01和P<0.05),但它们两者之间差异却不显著(P>0.05)。在体长上,CC基因型个体的体长显著长于CT基因型(P<0.05),TT基因型与CT基因型和CC基因型差异不显著(P>0.05)。在西农萨能奶山羊中,CC基因型在体高上则显著地低于CT和TT基因型(P<0.05),CT基因型和TT基因型之间差异不显著。不同基因型对体长和胸围不存在显著影响。
结果说明,IGF-IR基因R8基因座与西农萨能奶山羊的乳密度相关,R9基因座不同基因型对两个奶山羊品种的体尺性状存在显著或极显著的影响。
Using PCR-SSCP and DNA sequencing techniques, genetic variations of 14 loci of insulin-like growth factor-I (IGF-I) and its receptor (IGF-IR) genes were detected in 708 individuals of two Chinese dairy goat breeds (Guanzhong and Xinong Saanen). Their genetic structure and diversity were analyzed, too. Association analysis were carried out by using SPSS software(16.0) and Least Square Analysis method on milk production traits and body size traits at the candidate genes of the above-mentioned breeds. The objects were to discover the hereditary characteristics of these dairy goat breeds, to explore genetic markers with significant effects on economic important traits for efficient selection, and to provide genetic information for the dairy goat breeding program. The results were as follows:
1. Polymorphisms of IGF-I gene in two dairy goat breeds
The variations of five loci (P1~P5) in IGF-I gene were detected in Guanzhong and Xinong Saanen breeds. These loci contained exon1W, exon1, exon3, exon4, exon5 and their partial introns. In those five loci, only P1 and P4 loci demonstrated polymorphisms, locating in 5’- flanking region and intron 4, respectively. Compared with goat IGF-I gene sequence published in GenBank (D26119.2), the polymorphisms were caused by two novel mutations namely g.1617 G > A and g.5752 G > C.
Chi-square test results were as follows: in P1 locus, both Guanzhong and Xinong Saanen dairy goats were in Hardy-Weinberg equilibrium (P>0.05). Genetic diversity indexes (Ne/PIC/He) were 1.16/0.13/0.14 and 1.23/0.17/0.19 respectively. In P4 locus, Xinong Saanen dairy goat was in Hardy-Weinberg equilibrium (P>0.05), but Guanzhong breed was not (P<0.01). Genetic diversity indexes (Ne/PIC/He) were 1.43/0.26/0.3,1.72/0.33/0.42 for Guanzhon and Xinong Saanen. The result illustrated that, P4 locus owned abundant polymorphic information (0.25 2. Associations between genetic variation of IGF-I gene and milk production and body size traits in two dairy goat breeds
Two genotypes (AG and GG) were found at P1 locus in IGF-I gene. Genotype GG was the majority genotype. Using SPSS software (16.0) and Least Square Analysis method, the results showed that different genotypes at P1 locus did not have significant influence on the body size in both breeds and on milk yield at the first lactation and the second lactation in Xinong Saanen dairy goat breed. The six indexes of milk composition including fat, protein, TS, SNF, lactose and density were measured in this study. Xinong Saanen dairy goats with AG genotype were significantly higher than that with GG genotype in fat, but their density was significantly lower than that with GG genotype. There was no significant influence on the other four indexes.
In P4 locus, three genotypes, GG, CG and GG were detected. Genotype GG was the majority genotype in both breeds. About body size trait, Guanzhong dairy goats with GG genotype had larger chest circumference (P<0.05), and ones with CG were larger than CC (P<0.05). In Xinong Saanen breed, the body length of those dairy goats with CG was largest in all the three genotypes, significantly larger than CC and CG ones (P<0.05). In Xinong Saanen breed, individuals with CC genotype can produce higher milk yield than the other two genotypes (P<0.05) at the first and the second lactations, but the ones with CG genotype can get higher milk yield than the ones with GG at the second lactation (P<0.05). Different genotypes did not influence on the milk composition significantly.
The result illustrate that, different genotypes at P1 locus had association with milk fat and density in Xinong Saanen dairy goats; different genotypes at P4 locus can influence the body size and milk yield in dairy goats.
3. Polymorphisms of IGF-IR gene in two dairy goat breeds
Nine loci (R1~R9) in IGF-IR gene, which included exon 1 to exon 10 (without exon 7) and their partial introns were analyzed in 2 dairy goat breeds. Only two (R8 and R 9 loci) of them demonstrated the polymorphisms. Both the loci had three kinds of genotypes(CC, CT and TT). In locus P8, the C>T mutation in exon 9 was found for the first time (NC_007319: g26688 C>T). The mutation resulted in Leu 608 Leu in IGF-IR protein. The polymorphism in P9 locus was caused by T>C mutation in intron 10 (NC_007319: g28273 T>C).
Chi-square test results were as follows: R8 and R9 loci were not in Hardy-Weinberg equilibrium in the two breeds (P<0.01). Genetic diversity indexes of Ne/PIC/He in Guanzhong and Xinong Saanen dairy goats were 2.00/0.37/0.50, 1.97/0.37/0.50, respectively at P8 locus and 1.99/0.37/0.50, 1.95/0.37/0.50 respectively at R9 locus. The results illustrated that both the R8 and R9 loci owned abundant polymorphic information (0.25 4. Associations between genetic variation of IGF-IR gene and milk production and body size traits in two dairy goat breeds
Three genotypes (CC, CT and TT) were found at R8 locus in IGF-IR gene. The CT genotype was the majority genotype. But different genotypes at this locus did not have any significant influence on the body size in two breeds and on the milk yield at the first and the second lactations in Xinong Saanen breed (P>0.05). In the milk composition, Xinong Saanen dairy goats with CC genotype had heavier milk density than the CT ones (P<0.05), and the milk density difference between the individuals with TT genotype and CT one was not significant.
Three kinds of genotypes (CC, CT and TT) were detected at R9 locus, too. TT genotype was the majority one. Different genotypes at this locus did not have any significant influence on milk production traits in Xinong Saanen dairy goats. In Guanzhong breed, the body heights of individuals with CC genotype were greatly significantly higher than those with CT genotype (P<0.01), and the goats with TT genotype had higher body heights than CT one (P<0.05). The individuals with CC genotype were longer in body length than those with TT (P<0.05). In Xinong Saanen dairy goats, individuals with TT and CT genotypes had higher body heights than those with CC genotypes (P<0.05). Different genotypes did not influence significantly on the body length and chest circumference.
The result illustrate that, the polymorphism at P8 locus had association with the milk density in Xinong Saanen breed; different genotypes at P9 locus had great significant or significant influence on the body size in both breeds.
1. 2个奶山羊品种IGF-I基因多态性
本研究共检测了奶山羊IGF-I基因5个基因座(P1～P5),即外显子1W、1、3、4、5以及部分内含子区域。在所研究的5个基因座中,仅P1和P4基因座发现多态性,分别位于外显子1W和第4内含子上。与已公布的山羊IGF-I基因序列(GeneBank登录号:D26119.2)比对发现,该两处多态性是g.1617 G > A和g.5752 G > C点突变导致的,在羊上为首次发现。
卡方检验显示:在P1基因座上,关中奶山羊和西农萨能奶山羊均处于Hardy-Weinberg平衡状态( P>0.05 ) ,其群体遗传多态指数Ne/PIC/He分别为:1.16/0.13/0.14,1.23/0.17/0.19。在P4基因座上,西农萨能奶山羊处于Hardy-Weinberg平衡状态(P>0.05),而关中奶山羊则处于不平衡状态(P<0.01)。关中奶山羊和西农萨能奶山羊群体遗传多态指数Ne/PIC/He分别为:1.43/0.26/0.3,1.72/0.33/0.42。
上述结果表明,P4基因座在这2个奶山羊品种中(关中和西农萨能)多态信息含量中等丰富(0.25
在IGF-I基因P1基因座中检测出AG和GG 2种基因型,其中GG基因型在2个品种中都为优势基因型。利用SPSS(16.0)软件,运用最小二乘线性模型等方法分析发现,P1基因座不同基因型与2个奶山羊品种的体尺性状,西农萨能奶山羊第一胎和第二胎产奶量无显著相关(P>0.05)。在分析西农萨能奶山羊奶成分中的脂肪、蛋白质、干物质、非脂固形物、乳糖含量和乳密度6个指标中,AG基因型个体乳脂率显著高于GG型个体(P<0.05),而其乳密度则显著低于GG型个体(P<0.05),不同的基因型对其他4个指标无显著影响。
P4基因座上共检测出CC、CG和GG 3种基因型,其中GG基因型在2个奶山羊品种中是优势基因型。在体尺性状上,关中奶山羊GG基因型仅在胸围上显著大于CG型和CC型(P<0.05),而CG型又显著大于CC型(P<0.05);西农萨能奶山羊群体中,CG基因型个体的体长最大,显著大于CC型和GG型(P<0.05),GG型和CC型差异则不显著(P>0.05)。P4基因座不同基因型对于西农萨能奶山羊产奶性状的影响则表现为:CC基因型个体无论是在第一胎还是第二胎产奶量上都显著高于其他两种基因型(P<0.05),CG基因型个体在第二胎产奶量上又显著高于GG基因型(P<0.05),不同基因型对奶成分无显著影响。
上述结果表明,P1基因座不同基因型与西农萨能奶山羊奶成分中的乳脂率和乳密度存在相关,P4基因座不同基因型与奶山羊体尺性状和产奶量存在相关。
3. 2个奶山羊品种IGF-IR基因多态性
本试验共检测了奶山羊IGF-IR基因9个基因座(R1～R9),包含该基因1～10个外显子及部分内含子(除外显子7)。在这9个基因座上,仅在R8和R9基因座上发现了多态性,2个品种在这2个位点上都表现CC、CT、TT 3种基因型。首次在R8基因座外显子9上发现了C>T的突变(NC_007319:g26688C>T),该突变导致了IGF-IR基因编码的蛋白第608位上的Leu氨基酸发生了同义突变。而R9基因座上的多态性是由于第10内含子上T>C(NC_007319:g28273T>C)突变引起的。
卡方检验显示:在R8和R9基因座中,2个品种均处于Hardy-Weinberg不平衡状态(P<0.01)。在R8中,关中奶山羊和西农萨能奶山羊群体遗传多态指数Ne/PIC/He分别为2.00/0.37/0.50,1.97/0.37/0.50。在R9中则为1.99/0.37/0.50,1.95/0.37/0.50。说明,R8和R9基因座在2个奶山羊品种中多态信息含量为中等丰富(0.25
在IGF-IR基因R8基因座中检测出CC、CT、TT 3种基因型,其中CT型在2个品种中都为优势基因型。但是该基因座上不同的基因型对2个奶山羊品种的体尺性状和西农萨能奶山羊的第一胎、第二胎产奶量都不存在着影响(P>0.05)。在所检测的奶成分共6个指标中,CC基因型的西农萨能奶山羊的乳密度显著高于CT基因型(P<0.05),与TT型差异不显著(P>0.05),TT基因型和CT基因型之间差异也不显著(P>0.05)。
在R9基因座中同样检测出CC、CT、TT型3种基因型,其中TT基因型占多数。该基因座不同基因型对西农萨能奶山羊的产奶性状不存在着显著影响。在关中奶山羊中,虽然CC和TT基因型个体的体高分别极显著和显著的高于CT基因型(P<0.01和P<0.05),但它们两者之间差异却不显著(P>0.05)。在体长上,CC基因型个体的体长显著长于CT基因型(P<0.05),TT基因型与CT基因型和CC基因型差异不显著(P>0.05)。在西农萨能奶山羊中,CC基因型在体高上则显著地低于CT和TT基因型(P<0.05),CT基因型和TT基因型之间差异不显著。不同基因型对体长和胸围不存在显著影响。
结果说明,IGF-IR基因R8基因座与西农萨能奶山羊的乳密度相关,R9基因座不同基因型对两个奶山羊品种的体尺性状存在显著或极显著的影响。
Using PCR-SSCP and DNA sequencing techniques, genetic variations of 14 loci of insulin-like growth factor-I (IGF-I) and its receptor (IGF-IR) genes were detected in 708 individuals of two Chinese dairy goat breeds (Guanzhong and Xinong Saanen). Their genetic structure and diversity were analyzed, too. Association analysis were carried out by using SPSS software(16.0) and Least Square Analysis method on milk production traits and body size traits at the candidate genes of the above-mentioned breeds. The objects were to discover the hereditary characteristics of these dairy goat breeds, to explore genetic markers with significant effects on economic important traits for efficient selection, and to provide genetic information for the dairy goat breeding program. The results were as follows:
1. Polymorphisms of IGF-I gene in two dairy goat breeds
The variations of five loci (P1~P5) in IGF-I gene were detected in Guanzhong and Xinong Saanen breeds. These loci contained exon1W, exon1, exon3, exon4, exon5 and their partial introns. In those five loci, only P1 and P4 loci demonstrated polymorphisms, locating in 5’- flanking region and intron 4, respectively. Compared with goat IGF-I gene sequence published in GenBank (D26119.2), the polymorphisms were caused by two novel mutations namely g.1617 G > A and g.5752 G > C.
Chi-square test results were as follows: in P1 locus, both Guanzhong and Xinong Saanen dairy goats were in Hardy-Weinberg equilibrium (P>0.05). Genetic diversity indexes (Ne/PIC/He) were 1.16/0.13/0.14 and 1.23/0.17/0.19 respectively. In P4 locus, Xinong Saanen dairy goat was in Hardy-Weinberg equilibrium (P>0.05), but Guanzhong breed was not (P<0.01). Genetic diversity indexes (Ne/PIC/He) were 1.43/0.26/0.3,1.72/0.33/0.42 for Guanzhon and Xinong Saanen. The result illustrated that, P4 locus owned abundant polymorphic information (0.25
Two genotypes (AG and GG) were found at P1 locus in IGF-I gene. Genotype GG was the majority genotype. Using SPSS software (16.0) and Least Square Analysis method, the results showed that different genotypes at P1 locus did not have significant influence on the body size in both breeds and on milk yield at the first lactation and the second lactation in Xinong Saanen dairy goat breed. The six indexes of milk composition including fat, protein, TS, SNF, lactose and density were measured in this study. Xinong Saanen dairy goats with AG genotype were significantly higher than that with GG genotype in fat, but their density was significantly lower than that with GG genotype. There was no significant influence on the other four indexes.
In P4 locus, three genotypes, GG, CG and GG were detected. Genotype GG was the majority genotype in both breeds. About body size trait, Guanzhong dairy goats with GG genotype had larger chest circumference (P<0.05), and ones with CG were larger than CC (P<0.05). In Xinong Saanen breed, the body length of those dairy goats with CG was largest in all the three genotypes, significantly larger than CC and CG ones (P<0.05). In Xinong Saanen breed, individuals with CC genotype can produce higher milk yield than the other two genotypes (P<0.05) at the first and the second lactations, but the ones with CG genotype can get higher milk yield than the ones with GG at the second lactation (P<0.05). Different genotypes did not influence on the milk composition significantly.
The result illustrate that, different genotypes at P1 locus had association with milk fat and density in Xinong Saanen dairy goats; different genotypes at P4 locus can influence the body size and milk yield in dairy goats.
3. Polymorphisms of IGF-IR gene in two dairy goat breeds
Nine loci (R1~R9) in IGF-IR gene, which included exon 1 to exon 10 (without exon 7) and their partial introns were analyzed in 2 dairy goat breeds. Only two (R8 and R 9 loci) of them demonstrated the polymorphisms. Both the loci had three kinds of genotypes(CC, CT and TT). In locus P8, the C>T mutation in exon 9 was found for the first time (NC_007319: g26688 C>T). The mutation resulted in Leu 608 Leu in IGF-IR protein. The polymorphism in P9 locus was caused by T>C mutation in intron 10 (NC_007319: g28273 T>C).
Chi-square test results were as follows: R8 and R9 loci were not in Hardy-Weinberg equilibrium in the two breeds (P<0.01). Genetic diversity indexes of Ne/PIC/He in Guanzhong and Xinong Saanen dairy goats were 2.00/0.37/0.50, 1.97/0.37/0.50, respectively at P8 locus and 1.99/0.37/0.50, 1.95/0.37/0.50 respectively at R9 locus. The results illustrated that both the R8 and R9 loci owned abundant polymorphic information (0.25
Three genotypes (CC, CT and TT) were found at R8 locus in IGF-IR gene. The CT genotype was the majority genotype. But different genotypes at this locus did not have any significant influence on the body size in two breeds and on the milk yield at the first and the second lactations in Xinong Saanen breed (P>0.05). In the milk composition, Xinong Saanen dairy goats with CC genotype had heavier milk density than the CT ones (P<0.05), and the milk density difference between the individuals with TT genotype and CT one was not significant.
Three kinds of genotypes (CC, CT and TT) were detected at R9 locus, too. TT genotype was the majority one. Different genotypes at this locus did not have any significant influence on milk production traits in Xinong Saanen dairy goats. In Guanzhong breed, the body heights of individuals with CC genotype were greatly significantly higher than those with CT genotype (P<0.01), and the goats with TT genotype had higher body heights than CT one (P<0.05). The individuals with CC genotype were longer in body length than those with TT (P<0.05). In Xinong Saanen dairy goats, individuals with TT and CT genotypes had higher body heights than those with CC genotypes (P<0.05). Different genotypes did not influence significantly on the body length and chest circumference.
The result illustrate that, the polymorphism at P8 locus had association with the milk density in Xinong Saanen breed; different genotypes at P9 locus had great significant or significant influence on the body size in both breeds.
引文
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